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Recent tendencies in distribution of epiphytic bryophytes in urban areas: a Wrocław case study (south-west Poland). PDF

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Polish Botanical Journal 57(1): 231–241, 2012 RECENT TENDENCIES IN DISTRIBUTION OF EPIPHYTIC BRYOPHYTES IN URBAN AREAS: A WROCŁAW CASE STUDY (SOUTH-WEST POLAND) EWA FUDALI Abstract. The lists of moss and liverwort species found on tree-trunks in the Wrocław parks obtained in three recent surveys (in the years 2000, 2006 and 2011) were compared and changes in species composition and abundance of records were discussed to defi ne apparent recent trends in the bryo-epiphyte occurrence within the city. General ecological requirements of the species colonizing tree-trunks in relation to substratum reaction were analyzed. In the last decade there has been greater bryophyte colonization of tree-trunks in the city. This effort is realized by an infl ux of additional species from outside, mainly obligate epiphytes (both acidophilic and subneutrophilic), by expansion of urban epiphytes tolerant of city conditions - such as the acido- philic Dicranoweisia cirrata (Hedw.) Lindb. and eurybiontic Hypnum cupressiforme Hedw. and Amblystegium serpens (Hedw.) Schimp., as well as by species occurring in urban areas on other substrata (often on walls) but which are able also to colonize the bark of trees. Details of the epiphytes occurring in the Wrocław parks are here provided for the fi rst time. Key words: urban bryophytes, bryophyte dynamics, acidophilic epiphytes, Orthodicranum tauricum, Wrocław, Poland Ewa Fudali, Departament of Botany and Plant Ecology, Wrocław University of the Life and Environmental Sciences, Pl. Grun- waldzki 24A, 50-363 Wrocław, Poland; e-mail: [email protected] INTRODUCTION The second half of the 20th century is generally the preceding 30 years. Only two of them were regarded as a period of retreat and extinction of acidophilic. Among 24 obligate epiphytes recorded epiphytes due to increase in air pollution and high in the years 1992–1995 nine were not reported exposure of SO , NO and dust (for example, before the year 1950 (Bates et al. 1997). 2 x Barkmann 1958; Gilbert 1968, 1970, 1971; Rao During the 20th century many epiphytic mosses 1982). and liverworts almost completely disappeared from At the end of the 20th century Greven (1992) city centers and they remained in cities only in reported a new phenomenon, observed by him in parks, with species richness and frequency rather the forests of Holland, where there was an abun- low (Seaward 1979; Wittig 1991; During 1992; dant occurrence of epiphytic bryophyte species that Fudali 1994). In many cases only Hypnum cupres- had not been previously documented in the 1950s. siforme Hedw. and Dicranoweisia cirrata (Hedw.) They all showed a preference for acid substrata. Lindb. appeared to be sustainable in city condi- Greven (1992) named them ‘acidophilic epiphytes’ tions, although other epiphytic bryophytes were and their infl ux and range expansion resulted from noted sporadically within urban areas (Nordhorn- acidifi cation due to acid rain which altered the Richter & Düll 1982; Ron et al. 1987; Müller 1993; nature of the tree bark and thus increased the Fudali 1996, 2005; Vanderpoorten 1997; Solga number of habitats available for these epiphytes 1998; Hohenwalner 2000; Fojcik & Stebel 2001; to colonize. Surveys undertaken by Bates et al. Humer-Hochuimmer & Zechmeister 2001). (1997) across southern Britain showed an increase Müller (1993) analyzed the distribution of in the number of records of 13 obligate epiphytes mosses and liverworts within the town of Halle in comparison with their distributions mapped over in the 1990s and suggested the possibility of 232 POLISH BOTANICAL JOURNAL 57(1). 2012 bryo-epiphyte vegetation recovery in this town. was 583 mm), and quite frequent are series of This was subsequently confirmed by Richter wet years with an annual rainfall higher than et al. (2009) in a survey carried out in the years 600 mm. In the years 1997 and 1998, 2009 and 2005–2006. Recently, Stapper and Kricke (2004) 2010 summer fl oods appeared. The average an- reported a visible increase in the number of epi- nual temperature is 9°C; annual amplitude of phytic bryophytes found in the area of Dusseldorf temperature 19.2°C. Winters are short (65 days) town between 1979 and 2003. and mild, with frequent periods of warming in Considering the reports from German towns February up to 10–15°C. The most frequent winds and the fact that an expansion of acidophilic epi- are from the west (25% of days in a year) and phytes has been recently confi rmed from many south (23.1%). Within the center of Wrocław an regions of Poland (Stebel 2006; Stebel et al. 2008; urban island of warming was detected (Dubicka Fojcik 2011; Fudali et al. 2010) the author formu- & Szymanowski 2000). lated the following questions: have acidophilic The vegetation of the region is deciduous mes- epiphytes came to Wrocław town and established? ophilous forests with a prevalence of oaks and Are there any preferences in distribution? Have hornbeams (Matuszkiewicz et al. 1995) but the other bryophytes shown epiphytic tendencies in the surrounding area for many centuries has been used city in the last decade and what are their ecological for agriculture. In the last decade there has been requirements in relation to substratum reaction a strong increase of building investment within and type? the town. Preliminary results of studies on the bryophyte Parks chosen for study differ in their area dynamics in fi ve large parks of Wrocław (Fudali (from 2.52 ha to 1000 ha), history, architecture of 2007) revealed an increase in the number of species greenery and location in relation to the city center colonizing tree-trunks between 2000 and 2006 and (Fudali 2001) but the all are historical sites and a further infl ux of new epiphytes in 2011 (Fudali were established in the 19th century or at the begin- 2011). In the present paper a list of moss and liv- ning of the 20th century. Four of them are situated erwort species found on tree-trunks in 15 Wrocław within old historical center, others are situated out- parks in the years 2000, 2006 and 2011 with details side but still within built up areas (Fig. 1). of their occurrence, abundance and tree host pref- erences (published for the fi rst time) is presented MATERIAL AND METHODS and changes in the species composition, records frequency and distribution are discussed in relation Field studies entailed the making of a precise bryofl o- to recent trends in the bryo-epiphytes occurrence ristic inventory within 15 parks situated in the area of Wrocław town and the collection of herbarium speci- within the city. General ecological requirements mens for taxonomic identifi cation. In each park every in relation to substratum reaction of the species tree was examined. For each occurrence of a bryophyte colonizing tree-trunks were analyzed to determine species the tree host was identifi ed and height of a bryo- the number of the acidophilic epiphytes. phyte patch situated on the tree-trunk (up to 50 cm or Wrocław is one of the biggest towns in Poland higher) recorded. The studies were regularly repeated and occupies an area of 290 km² with a population at 5–6 years intervals since 2000. Since 2006 some ad- of almost 700 000 inhabitants. It was established ditional observations have been made along main streets at the end of the 12th century and was in the past and within 10 selected housing estates with green af- an administrative, political and industrial center forested areas. In total 1680 records were made and 136 herbarium in the region of Lower Silesia. Through the town, specimens were collected. The latter are deposited in the from SE to NW, fl ows the river Odra with its four herbarium [KRAM-B]. Nomenclature of mosses follows tributaries. Ochyra et al. (2003) with the exception of Rosulabryum The annual rainfall is highly variable and moravicum (Podp.) Ochyra & Stebel. General ecological ranges between 318 to 892 mm annually (av- requirements of the species analyzed in relation to sub- erage annual precipitation in the 20th century strate reaction were extracted from Dierssen’s (2001) E. FUDALI: RECENT TENDENCIES IN DISTRIBUTION OF EPIPHYTIC BRYOPHYTES IN URBAN AREAS 233 Fig. 1. Situation of the parks studied on the background of the Wrocław town land use (according to Dubicka & Szymanowski 2000, modifi ed). 1 – parks studied, 2 – densely built up area, 3 – industrial buildings, 4 – housing settlements, 5 – municipal forests, 6 – open area, rural or neglected; C – historical center of the town. characterization of the European bryophytes ecological cies were recorded (Table 1), often growing at amplitudes. a height of 0.5 to 3.0 m above the ground. Among them only Orthotrichum diaphanum Schrad. ex Brid., and Ceratodon purpureus (Hedw.) Brid., RESULTS were frequently recorded and abundant. SPECIES DIVERSITY OF COLONIZING BRYOPHYTES The bryophyte species were collected from 26 tree species, but the most frequently colonized tree Altogether 36 species were recorded from tree host species were: Quercus robur L. (24 species), trunks in urban parks, along major streets, and Fraxinus excelsior L. (18), Acer platanoides L. within housing estates, including two liverworts (17), cultivars of Populus L. (17) and Betula pen- (Lophocolea heterophylla (Schrad.) Dumort. and dula Roth. (16). Ptilidium pulcherrimum (Weber) Vain.) and 34 Trunks of trees growing individually along mosses (Table 1). In the parks, most often the main streets were devoid of bryophytes. species colonized the lower parts of tree trunks. Only Dicranoweisia cirrata and Hypnum cupressi- INCREASE SPECIES RICHNESS AND FLUCTUATIONS forme occurred also quite abundantly on tree trunks IN SPECIES COMPOSITION higher than 50 cm above the ground. Bryophytes were also recorded on a few trees occurring within Since the year 2000 the number of species colo- some of housing estates – in total four moss spe- nizing tree trunks in Wrocław’s parks has increased 234 POLISH BOTANICAL JOURNAL 57(1). 2012 Table 1. List of species colonizing tree-trunks in the Wrocław’s parks in the period of 2000–2011 ordered according to their maintenance and the year of their record with brief characteristics of their station’s abundance and range. A – tree trunk at height more than 50 cm above the ground level, B – tree trunk up to 50 cm above the ground level, N – total number of records, O – total number of parks, in which species was noted, PC – parks situated in the city center, PP – parks out of the center; ¹ – in Wrocław Dicranum scoparium occurred exclusively on trees (Fudali 2001, 2007, 2011), * – species noted also on few trees within housing estates studied, ● – specialized epiphytic species, • – epiphytic-epilithic species; abbreviations for tree species: Ac – Acer platanoides L., Ae – Aesculus hippocastanum L., Al – Alnus glutinosa (L.) Gaertn., An – Acer negundo L., Ap – Acer pseudoplatanus L., Be – Betula pendula Roth., Ca – Carpinus betulus L., Co – Cornus mas L., Cr – Crategus L. sp., Cs – Cerasus avium (L.) Moenck, ×Fr – Fraxinus excelsior L., Pa – Picea abies (L.) H. Karnst, Pi – Pyrus communis L., Pl – Platanus acerifolia (Aiton) Willd., Pr – Prunus L. sp., Ps – Pinus sylvestris L., Px – cultivars of Populus L. sp., Qu – Quercus robur L., Qr – Quercus rubra L., Ro – Robinia pseudoacacia L., Sa – Salix alba L., Th – Thuja occidentalis L., Ti – Tilia cordata Mill., Ul – Ulmus laevis Pall., Tulip – Liriodendron tulipifera L. 2000 20062011SpeciesPPNOPPNNOPCPPNO 2000-2011SPECIESPERSISTENTINTHEYEARSA11752281-Ac 2-Ac,Fr; 1-Ti,Px,Qu1-Px10-Ac; 4-Cs; 3-Fr; 2-Ro; 1-Qu,Sa*Amblystegium serpens 45-Ac; 26-Qu; 20-Fa; 18-Fr; 13-Qr; 5-Ac,Fr; 3-Qu; 11-Fr; 9-Ac; 4-Qr; (Hedw.) Schimp.1889B1662861-Ac9-Ca,Px; 8-Al,Ro,Sa; 7-Ti; 4-Be,Pa; 1-Fa,Ro,Ti2-Ro,Qu,Pr; 3-Ul; 1-Ae,Ap,PlA 0032 332-Be; 1-Ac2-Fr;1-PxBrachytheciastrum velutinum 4-Qr; 2-Fr,Qu; 18-Ac; 9-Be; 5-Fr; 4-Al,Qr,Qu,Ro; (Hedw.) Ignatov & HuttunenB12124 6071-Ac,Qu1-Ac,Be,Ro,Ti2-Ca,Cr,Ti,Ul; 1-Ap,Sa,Th,PlA 0 0 221-Be,Sa*Brachythecium rutabulum 17-Qu; 14-Ac; 7-Be; 4-Fr; (Hedw.) Schimp.656B1141 ,Ro,Fa; 2-Ca; 1-Cs2-Ro,Ti3-Al,Cr,Qr1-Ae,Ap,Pa,Px,Th,TiA21431272-Be3-Qu; 1-Ro,Ti1-Ac,Ae,Ro3-Ac; 1-Be,Fr,Qu,Px,Ro,Sa*Ceratodon purpureus Hedw.2-Al,Fr,Ro; 12-Ac; 6-Qu; 5-Fr,Qr,Ro; 3-Fa; 2-Be; B32944292-Fr; 1-Ac1-Px1-Ac,Be,Qr1-Ca,Th,Ul18-Ac; 16-Be; 10-Qu; 5-Px; 3-Al,Cs,Ro; A32103 6453-Qu5-Qu; 2-Ac,Be; 1-Fr2-Ac,Qr; 1-Ca,Cr,Fa,FrDicranoweisia cirrata (Hedw.) ●Lindb.48-Qu; 18-Be; 13-Al; 8-Ac; 7-Ti; 5-Px; B64104 11352-Ac,Qu; 1-Be,Pa4-Be,Qu; 1-Fr,Ro4-Qr; 3-Pa; 2-Ae,Fr; 1-Fa,Ro,Pi 41 22A11,Qu1-Px,Qx1-Qu2-Px; 1-Be¹Dicranum scoparium Hedw.●0 B 0 422-Qu; 1-Be,Fr9-Be,Qu; 5-Px; 19-Qu; 15-Ac; 7-Px; 6-Fr; 5-Be,Cs,Sa; A9432678104-Px; 2-Qu,Ro; 1-Ca1-Ac,Px4-Ac,Fr; 1-Ro,Ti4-Al,Ro; 3-Qr; 1-Cr,Ti,UlHypnum cupressiforme Hedw.167-Qu; 58-Ac; 36-Be;32-Al; 26-Fr; 31-Qu; 13-Ac; 31-Qu; 5-Ac; 4-Px; 6474209B446Fr; 3-Ti; 1-Ac24-Qr; 17-Px,Ti; 8-Ca; 7-Fa,Ro; 6-Sa; 5-Be,3-Fr; 2-Be; 1-Ro2-Cr,Px,Ro; 1-Ca 5-Pa; 3-Ae,Ul; 1-Ae,Ap,Pi,TulipA1111 001-Qu1-BeLophocolea heterophylla (Schrad.) Dumort.B5211 1024-Qu;1-Pa1-Qu7-Qu; 1-Be,Pl,Ti E. FUDALI: RECENT TENDENCIES IN DISTRIBUTION OF EPIPHYTIC BRYOPHYTES IN URBAN AREAS 235 0 1 2 1 2 2 3 1 3 5 1 3 3 6 6 86 1 1 1 12 1 212557 0 1 3 1 3 17 3 1 5 24 2 5 6 18 26 1738 1 2 1 16 1 21391921 Ul a, 1-Qr 2-Qu; 1-Be 1-Qu 3-Qu 9-Be; 3-Qu; 2-Al; 1-Fa,Fr,Px 1-An,Cs,Px 1-Sa 2-Ac,Px; 1-Ro10-Fr; 5-Ac; 3-Px; 2-Qr; 1-Ae,Be,Qu,Ro,Sa 2-Qu 3-Cs,Px,Sa 2-Ac,Qu; 1-Cs,Th 7-Ac; 4-Cs; 2-Fr; 1-Ap,Cr,Qu 13-Ac; 5-Qu; 3-Px; 2-Fr; 1-Al,Be 5-Ac; 4-Qu; 2-Fr; 1-Al,Ap,Be,Cs,Ti22-Qu; 6-Ac; 2-Be,Px; 1-Al,Fr,Pl,S 1-Ac 2-Ac 1-Qu 1-Qu4-Qu; 1-Be,Fr 1-Sa 1-Ac,Qu1-Qu2-Ac; 1-Cs1-Qu,Px,Sa7-Px; 3-Ac,Fr; 1-Ap,Cr,Fr 5-Ac; 3-Fr,Px; 2-Ro; 1-Sa cu AQ c,Px x ce c c,Frx; 1-r; 1- A P AB A APF 1- 1- 1-1- 1- 3-2-6- 0 1 2 1 2 3 2 1 1 1 1 1 0 0 2 03 0 0 0 00 0 000000 0 1 5 1 4 3 2 1 2 3 1 1 0 0 3 03 0 0 0 00 0 000000 Ti Be, Pa Ro Qu Fr 1-Qu 3-Qu; 1- 1-Ro 4-Qu 1-Be,Qu, 1-Fr,Px 1-Fr 2-Qu 1-Px,Qu, 2011GIN 1-Qu 1-Px 1-Cs,Fr, 2-Qu; 1- N 1 0 1 1 1 1 1 1 0 2 RI 0 0 0 0 0 00 0 0 0 00 0 000000 R U 2 0 2 1 2 2 1 1 0 5 C 0 0 0 0 0 00 0 0 0 00 0 000000 C O L L TI S o D R N c, A Qu,Ro Pa,Qu Pa Qu,Ti Qu Px Qu Qu; 1-A 2006 N 2011N 1- 1- 1- 1- 2- 1- 1- 3- EI EI M M A B B B B B A B A B TI B A B A B AB TI B B B AB B ABABAB T T Orthodicranum montanum (Hedw.) Loeske● Plagiothecium curvifolium Schlieph. ex Limpr.Plagiothecium denticulatum (Hedw.) Schimp.Plagiothecium laetum Schimp.●Pohlia nutans (Hedw.) Lindb. Pylaisia polyantha (Hedw.) Schimp.• Rosulabryum moravicum (Podp.) Ochyra & Stebel• S PECIESREPORTEDFORTHEFIRS Mnium hornum Hedw. Orthotrichum affi ne Schrad. ex Brid.● Orthotrichum pallens Bruch ex Brid.• Platygyrium repens (Brid.) Schimp.● S PECIESREPORTEDFORTHEFIRS Bryum argenteum Hedw. Dicranella heteromalla (Hedw.) Schimp. Herzogiella seligeri (Brid.) Z. Iwats. Hypnum pallescens (Hedw.) P. Beauv.● Leptodictyum riparium (Hedw.) Warnst. Orthodicranum tauricum (Sapjegin) Smirnova● Orthotrichum anomalum Hedw. *Orthotrichum diaphanum Schrad. ex Brid.• 236 POLISH BOTANICAL JOURNAL 57(1). 2012 O 1 5 3 1 1 1 1 1 1 00 0 0 N 1 7 7 2 1 1 1 2 1 00 0 0 1 01 Ti 2 r,Sa, 1-Ti Ac,F Qu; x; 1- c,Fr, e,Ro x c c,Be PP Ac 2-P 2-A 1-B 1-P 1-A 1-A x c x C P A P P 1- 1- 1- O 0 0 0 0 0 0 0 0 0 01 1 1 N N 0 0 0 0 0 0 0 0 0 01 1 1 6 0 0 2 PP 1-Fr 2011 1-Fr 1-Qu N O 0 0 0 0 0 0 0 0 0 10 DI 0 0 1 N 1 U N 0 0 0 0 0 0 0 0 0 0 10 O 0 0 2 F 2000 OUNDIN DNOTRE REF AN OT 06 PP 6 ANDN 1-Qu 20MEIN A A B B A B A B A 200 AB TTI B A Table 1. Continued. Species Orthotrichum pumilum Sw. ex anon. • Orthotrichum speciosum Nees● Plagiomnium cuspidatum (Hedw.) T. Kop. Sanionia uncinata (Hedw.) Loeske• Sciuro-hypnum populeum (Hedw.) Ignatov & Huttunen Syntrichia virescens (De Not.) Ochyra• S 2000 PECIESREPORTEDINAND Leskea polycarpa Hedw.● SPECIESREPORTEDFORTHEFIRS Aulacomnium androgynum (Hedw.) Schwägr. Ptilidium pulcherrimum (Weber) Vain. E. FUDALI: RECENT TENDENCIES IN DISTRIBUTION OF EPIPHYTIC BRYOPHYTES IN URBAN AREAS 237 Table 2. Number of species reported from tree trunks in the years of the bryofl oristical surveys. A – tree-trunks at height more than 50 cm above the ground level, B – tree-trunks at height up to 50 cm above the ground level. 2000 2006 2011 Category of parks A B A B A B Number of species 16 22 33 All the parks 10 13 11 20 21 32 0 1 10 Parks in the town center 0 0 0 1 9 7 16 21 32 Parks situated out of the center 10 13 11 19 21 30 Fig. 2. Substratum preferences of species recorded from tree markedly, especially in parks situated out of the trunks in the parks with regards to the date of their records. historical center (Table 2). Comparison of the Species occurring on: G – soil (exclusively), E – trees bark species lists obtained in the years of observation (exclusively), L – walls (exclusively), X – decayed wood (ex- clusively), P – more than two substratum types. shows that among 36 species reported 15 (42 %) have maintained a presence within the town’s parks during the 11 years and only one species, Leskea polycarpa Hedw., recorded in 2000 and 2006 has not been found subsequently (Table 1). In 2006, 7 species were recorded for the fi rst time, but two of them, Aulacomnium androgynum (Hedw.) Schwägr., and Ptilidium pulcherrimum, were not refound in 2011. In 2011 an additional 14 species were collected from tree trunks for the fi rst time. ECOLOGICAL CHARACTER OF THE SPECIES RECORDED FROM TREE-TRUNKS Analysis of the species’ habitat reaction and sub- Fig. 3. General ecological requirements in relation to the sub- stratum preferences shows that only 10 species stratum reaction of the species recorded from tree trunks in are obligate epiphytes (Fig. 2). Among others the the parks with regards to the date of their records. Reaction of most frequent are 7 species occurring both on tree substratum: AC – acid, BA – basic, c – considerably, h – highly, bark and walls (epilithic-epiphytic species) and m – moderately, SB – subneutral. 11 species colonizing more than two substratum considerably acid to subneutral – 31%. The per- types (multi-substrata species). The latter dominate centage incidence of subneutral and subneutral among the persistent species while those noted for to basophilic species is also high – 25% (Fig. 3). the fi rst time in the years 2006 and 2011 show Most of the latter species have appeared recently ecological differentiation and more than half of (reported for the fi rst time in the year 2011). them (11 species or 52%) were obligate epiphytes or epiphytic-epilithic species. CHANGES IN SPECIES ABUNDANCE Analysis of the species requirements towards AND DISTRIBUTION substratum reaction revealed a high percentage incidence of both highly to moderate acidophilic The 7 (47%) ‘persistent’ species have shown species – 33% and eurybiontic species occurring constant and multiple increase of the number of in habitats of wide amplitude of reaction from records and parks in which they were observed 238 POLISH BOTANICAL JOURNAL 57(1). 2012 (Table 1). These are: multi-substratum species The changes described above in species abun- of wide amplitude in relation to the substratum dance and distribution refer to parks situated out reaction (from considerably acid to subneutral) of the historical town center. Trees situated in as Amblystegium serpens, Brachytheciastrum parks within the city center have been sporadi- velutinum (Hedw.) Ignatov & Huttunen, Brach- cally colonized by mosses (Table 1). However ythecium rutabulum (Hedw.) Schimp., Ceratodon the bryo-fl oristic inventory done in the year 2011 purpureus and Hypnum cupressiforme, acidophilic revealed a striking increase in the number of spe- epiphyte Dicranoweisia cirrata and epiphytic-epi- cies colonizing tree-trunks in the period between lithic species of undetermined reaction preferences 2006 and 2011 (10 new species). They show high (Dierssen 2001) Rosulabryum moravicum (Podp.) ecological similarity (Table 3). Ochyra & Stebel. The remaining ‘persistent’ spe- cies have not shown such spectacular changes in DISCUSSION AND CONCLUSIONS their abundance and distribution. Three of the species reported for the first During the last 11 years the number of species time in the year 2006 also have shown constant colonizing tree trunks in the Wrocław municipal tendency to establishment (Table 1). These are parks doubled from 16 to 33, including 5 new two obligate epiphytes of quite wide ecological obligate epiphytic species not previously noted in amplitude in relation to the substratum reaction the city (Fudali 2001): Orthodicranum tauricum (from moderately acid to subneutral) Orthotrichum (Sapjegin) Smirnova, Hypnum pallescens, Orthot- pallens Bruch ex Brid., and Platygyrium repens richum affi ne, O. pumilum Sw., O. speciosum. Only (Brid.) Schimp., as well as the epiphytic-epilithic one eiphytic species, Leskea polycarpa, was not species Orthotrichum pallens preferring habitats refound in 2011. of subneutral reaction. These new epiphytes differ in their require- Most of the species recorded for the fi rst time ments towards substratum reaction: three of them in the year 2011 occurred sporadically with the prefer subneutral to basic habitats (species of the exception of subneutrophilic species: the epiphyte genus Orthotrichum) and two are acidophytes Orthotrichum speciosum Nees and epiphytic-epi- (Orthodicranum tauricum, Hypnum pallescens). lithic species O. diaphanum as well as epilithic The surveys demonstrate that some acidophilic O. anomalum Hedw., colonizing moderately acid epiphytes sensu Greven (1992) have reached to subneutral habitats, which were noted several Wrocław town recently and are establishing. They times. were reported from the largest parks situated out of the city center and their sites were in every Table 3. Ecological character of the species recorded in 2011 case situated on the parks’ borders in the vicinity on tree-trunks in the parks situated in the city center. Explana- of a street. Quercus robur trunks seem to be a fa- tions as for Figs 2 and 3. voured substrate. Orthodicranum tauricum is considered to be Substratum Substratum Name of species a species signifi cantly expanding its range in reaction preferences Europe in the second half of the 20th century Amblystegium serpens c AC-SB P Bryum argenteum SB G-L (Söderstrom 1992). In Poland, until 1986 the spe- Ceratodon purpureus m AC-SB P cies was known only from eight localities but Hypnum cupressiforme h AC-SB P since the late 1980’s the number of its new sta- Orthotrichum diaphanum SB-BA E-L tions increased remarkably to about 70 (Stebel Orthotrichum pallens SB E-L et al. 2008). Some of these stations are situated Orthotrichum pumilum SB-BA E-L within administrative borders of Polish cities Orthotrichum speciosum SB E and towns (Katowice, Kraków, Łódź, Poznań, Platygyrium repens m AC-SB E Rybnik, Zgierz) but always in municipal forests Syntrichia virescens SB-BA E-L or nature reserves. In Wrocław the species was E. FUDALI: RECENT TENDENCIES IN DISTRIBUTION OF EPIPHYTIC BRYOPHYTES IN URBAN AREAS 239 recorded in three distant sites within two large dorf (Stapper & Kricke 2004) and Halle (Richter historical parks (Zachodni, Szczytnicki) estab- et al. 2009). lished in the past within natural forests but pres- A number of other normally epilithic species ently surrounded by buildings. Orthodicranum are occurring with remarkable frequency as epi- tauricum was also reported from the German city phytes on tree trunks (Fudali 2005): Dryptodon Düsseldorf (Stapper & Kricke 2004) as very rare, pulvinatus (Hedw.) Brid., and Bryum argenteum but without details of its habitat. The reasons for Hedw., in Düsseldorf (Stapper & Kricke 2004), the spread of Orthodicranum tauricum are still Orthotrichum pumilum in Halle (Richter et al. unclear; some authors suggested that it may be 2009) and Orthotrichum pallens in Wrocław. associated with acidifi cation of habitats (Greven The appearance of epiphytic tendencies in spe- 1992; Söderstrom 1992) or climatic changes cies overgrowing rocks was reported already by (Stebel et al. 2008). Barkmann (1958). Considering the problem of the supposed ex- The recent increase in the number of species pansion of acidophilic epiphytes in urban areas it colonizing tree trunks in towns has been earlier is worth stressing that Dicranoweisia cirrata, clas- reported from Germany. Surveys undertaken by sifi ed by Greven (1992) in this ecological group, Stapper & Kricke (2004) in the Düsseldorf town showed a remarkable increase in a number of sta- reported signifi cant increase in corticolous bryo- tions in the Wrocław parks. Similar changes in this phyte species diversity between 1979 and 2003. On species distribution and frequency were reported the trunks of 1015 trees they found 23 bryophyte from two German towns: Düsseldorf (Stapper species of which 11 are also reported in Wrocław. & Kricke 2004) and Halle (Richter et al. 2009). They identifi ed three species of Orthotrichum, of The species is known to be tolerant of acid pre- which O. affi ne and O. diaphanum occurred with cipitation (Dierssen 2001). high frequency. The studies carried out in Wrocław parks Richter et al. (2009) encountered 40 bryophyte revealed also a remarkable increase in the spe- species colonizing tree trunks within Halle town cies richness of bryophytes preferring substrata in the years 2005–2006, while in the same area of subneutral or basic reaction, mainly rocks Müller (1993) collected from trees 21 species, in- (or walls in towns) and tree bark, including 4 cluding 10 obligate epiphytes. Unfortunately these species of Orthotrichum genus. These species authors did not provide a full list of recently re- showed a tendency to colonize tree bark even in corded species but they listed two new epiphytes: parks situated in the city center and those trees Orthotrichum affi ne and Ulota bruchii Hornsch. growing within housing estates. The Orthotri- Except for Pylaisia polyantha all species noted by chum species were collected mainly from the Müller (1993) still occurred. They also noticed that trunks of Fraxinus excelsior, hybrids of Populus some obligate epiphytic species, especially Orthot- sp., Salix alba L. and Acer platanoides having richum, showed a strong increase in the number bark that is known to be slightly acid (Bark- of sites and population abundance but only within mann 1958). that part of Halle town situated on the fl oodplain The most spectacular increase occurred with of the Saara river. The 18 species reported from Orthotrichum diaphanum, commonly occurring tree trunks in Halle (Müller 1993) occurred also in Wrocław on walls and concrete (Fudali 2001). on trees in the Wrocław parks. It seems probable that a strong intensifi cation of The species of bryo-epiphytes reported re- building investment within the town in the last cently from tree trunks within the cited German decade has promoted the expansion of subneutral towns seem to be heterogenic with regard to sub- epilithic-epiphytic species and their settlement on strate specialization. The list contains not only tree bark covered with building dust containing obligate but facultative epiphytes as well, such concrete. On tree trunks Orthotrichum diaphanum as epiphytic-epilithic species, epixylic species, was noted with high frequency also in Düssel- typical epiliths, species colonizing mainly humus 240 POLISH BOTANICAL JOURNAL 57(1). 2012 and multi-substratum species. This is also observed REFERENCES in the parks of Wrocław. In all towns compared some species showed BARKMANN J. J. 1958. Phytosociology and ecology of cryp- a clear increase in frequency and colony spread. togamic epiphytes. Van Gorcum & Comp. N. V. – G. H. Hak & Dr H. J. Prakke, Assen. These are: Dicranoweisia cirrata, as well as the eurybiontic multi-substratum species Hypnum cu- BATES J. W., PROCTOR M. C. F., PRESTON C. D., HODGETTS N. G. & PERRY A. R. 1997. Occurrence of epiphytic bryo- pressiforme and the subneutral epilithic-epiphytic phytes in a „tetrad” transect across southern Britain. 1. species Orthotrichum diaphanum. A similar ten- Geographical trends in abundance and evidence of recent dency, but only in Halle and Wrocław, was shown change. J. Bryol. 19: 685–714. also for the subneutral epilithic-epiphytic species DIERSSEN K. 2001. Distribution, ecological amplitude and phy- Orthotrichum pumilum, the acidophilic Dicranum tosociological characterization of European bryophytes. Bryophyt. Biblioth. 56: 1–289 scoparium Hedw., which in these cities occurred exclusively on trees, and the obligate epiphyte DUBICKA M. & SZYMANOWSKI M. 2000. Struktura miejskiej wyspy ciepła i jej związek z warunkami pogodowymi Platygyrium repens of wide amplitude of sites i klimatycznymi Wrocławia. Acta Univ. Wratislav. Studia reaction. Geografi czne 74: 99–118. As every survey referred to here has utilized DURING H. J. 1992. Endangered bryophytes in Europe. Tree different methods in relation to the area studied and 7(8): 253–255. the trees investigated, the results are not strictly FOJCIK B. 2011. Mosses of the Cracow-Częstochowa Upland comparable. However, in general they confi rm in relation to anthropogenic transformations of the plant Müller’s (1993) suggestion of bryophyte recovery cover. Prace Naukowe Uniwersytetu Śląskiego 2800: within towns and allow the conclusion that in the 1–232 (in Polish with English summary). last two decades bryophytes have started to recolo- FOJCIK B. & STEBEL A. 2001. Ecological and spatial structure of bryofl ora of Katowice town. Centrum Dziedzictwa Przy- nize trees in towns. This has occurred through the rody Górnego Śląska. Materiały, Opracowania 5: 1–128, infl ux of new species from outside, mainly obligate (in Polish with English summary). epiphytes (both acidophilic and subneutrophilic), FUDALI E. 1994. Species diversity and spatial distribution of by expansion of existing urban epiphytes such as bryophytes in urban areas – a case study of the city of the acidophilic Dicranoweisia cirrata and eury- Szczecin. Fragm. Florist. Geobot. 39(2): 563–570. biontic Hypnum cupressiforme and Amblystegium FUDALI E. 1996. Distribution of bryophytes in various urban- serpens, that appear to be sustainable in city condi- use complexes in Szczecin. Fragm. Florist. Geobot. 41: tions as well as by the species occurring in towns 717–745. on other substrata (often on walls). FUDALI E. 2001. The bryophytes of the town parks and cem- In general, a share of acidophilic epiphytes is eteries of Wrocław (W Poland). Przegląd Przyrodniczy 12(1–2): 3–20 (in Polish with English summary). less signifi cant than the participation of eurybiontic species tolerating habitats of wide substrate reac- FUDALI E. 2005. Bryophyte species diversity and ecology in the parks and cemeteries of selected Polish cities. Wydaw- tion amplitude from considerably or moderately nictwo Akademii Rolniczej, Wrocław. acid to subneutral, and an incidence of species FUDALI E. 2007. Bryophyte dynamic tendences in urban parks preferring habitats of subneutral or basic reaction. – a case study of Wrocław . Annales Silesiae 35: 11–20 The latter are mainly epilithic-epiphytic species, (in Polish with English summary). with a remarkable representation of the genus Or- FUDALI E. 2011. Contemporary changes in urban bryophytes: thotrichum. a case studyof Wroclaw parks carried out in 2000, 2006 and 2011. Acta Botanica Silesiaca 6: 63–77 (in Polish ACKNOWLEDGEMENTS. I thank Dr. Vítěslav Plàšek with English summary). (University of Ostrava) who confi rmed my determi- FUDALI E., RUSIŃSKA A. & RUTKOWSKI P. 2010. Dynamic nation of Orthotrichum pallens specimens and for tendencies in the bryofl ora of the nature reserve “Bukowe determining the specimens of O. pumilum, Professor Zdroje” (Puszcza Bukowa Forest near Szczecin) in the Rodney D. Seppelt (Australian Antarctic Division) for years 1969–2006. Botanica – Steciana 14: 3–10 checking the English, and the anonymous reviewers GILBERT O. L. 1968. Bryophytes as indicators of air pollution for helpful comments. in the Tyne Valley. New Phytol. 67: 15–30.

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